science and natural history

PRE-BLIGHT ABUNDANCE OF AMERICAN CHESTNUT IN

by C harles C. Rhoades, Department of Forestry, University of Kentucky, and Clare Park, Natural Resources Program, University of Kentucky

INTRODUCTION he Commonwealth ofKentucky extends west from the Appalachian T Mountains to the Mississippi River, spanning a variety of physio­ graphic regions (Fenneman, 1938) and forest types (Braun, 1950; Smalley, 1986). American chestnut (Castanea dentata) was a major component of both mixed mesophytic and western mesophytic forests, the two dom­ inant forest types in Kentucky (Braun, 1950). With the exception of the limestone-dominated in central Kentucky and portions of the western extreme of the state, chestnut was found throughout Kentucky (Saucier, 1973; Russell, 1987). The chestnut blight fungus ( Cryphonectria parasitica) spread across the southern Appalachians and Kentucky during the 1920's and 1930's (Cochran, 1990). By the mid- to late- 1930's, the blight had spread throughout the state; by the mid-1950's, the American chestnut as an overstory tree was virtually extinct. Forest pathologists and geneticists from various institutions currently predict that blight-resistant seedlings will be available for distribution and outplanting within five to fifteen years (Hebard, et al. 2000; S. Anagnostakis, Conn. Ag. Expt. Station, pers comm.). These predictions have spawned public interest in reintroduc­ tion ofAmerican chestnut to eastern forest ecosystems. In Kentucky, where production of hardwood lumber is a major source of financial gain, remrn of the chestnut combines nostalgia for the fo rests of the Appalachian fo re­ bearers, tl1e restoration of original fo rest species composition and forest dynamics, and the potential of regional economic de ..elopme nt. Pollen analysis has confirmed the existence of American chestnut in Kentucky's forests for the past several millennia and has linked increased chestnut density with Native American burning and clearing practices (Delcourt and Delcourt, 1997 & 1998). While range maps detail tl1e pre­ blight distribution ofAmerican chestnut (Saucier, 1973; Russell, 1987),

36 JOURNAL OF THE AMERIC.AN CHESTNUT FOUNDATION science and natural history very little quantitative information exists regarding the tree's abundance within Kentucky forests (Braun, 1935 ). Such information will help focus and prioritize Kentucky's chestnut reintroduction activities. The objective of this project was to survey information regarding the historic distribution and abundance of chestnut across Kenrucky. We reviewed independent sources of historic information that each provid­ ed county-level estimates of chestnut canopy cover, relative stem densi­ ty, or timber volume. The sources included land deeds, USDA chestnut blight survey notes, and a statewide inventory of standing timber. The data sources corresponded to the 30-year period leading up to the onset ofchestnut blight disease in Kentucky. Taken separately, the inherent bias­ es of historic data may weaken their ability to describe past conditions (Whitney and DeCant, 2001). Comparison between several indepen­ dent sources, however, should strengthen their predictive value.

SOURCES OF HISTORIC INFORMATION The statewide distribution and abundance ofAmerican chestnut was esti­ mated from an inventory of standing timber resources produced in 1919 by the USDA, Forest Service, and the Kentucky Department of Geology and Forestry (Barton, 1919). This inventory ranked the board footage of the dominant forest species for the 119 Kentucky counties established at that time. One county (McCreary) was established following the data collection phase of the project. Between 7 and 18 tree species were recorded for each county. County area, forested area, and the average stand volume were also estimated. While we have been unable to locate details regarding the data collection protocol, this record provides a unique "snap-shot" of the historic importance of chestnut as a timber species at the time of the arrival of blight in Kentucky. In Kentucky, land deed surveys were historically delineated by corner trees or distinct topographic features (i.e. streambanks, exposed rock out­ crops). Individual land deeds provide a species tally based on the corner trees, along \vi th the ownership, size, and location of the plot (Whitney and DcCant, 200 l ). Individual deeds were registered and archived at county land offices across the Commonwealth from the time ofs tatehood in 1792, through the early part ofthe 20th century. Although some of these records have been destroyed by fire, most are still available. Each land survey pro­ \~des a unique sample of tree species composition within a specific county

VOLUME XV, NUMBER 1 • SUMMER/FALL 2001 37 science and natura l history

and time period. For parcels deeded around the rum of the 20th century ( 1890-1910), we tallied the corner tree species on a minimum offive deeds per cow1ty for about half the counties in Kentucky (n = 54). As the chestnut blight disease radiated south and west from its point of introduction in New York City through the eastern deciduous forest, plant pathologists from the US Department of Agriculture tracked its progress. Between 1924 and 1931 USDA field agents traveled through Kentucky surveying the abundance and health of chestnut stands. Their systematic assessment noted local topography and overall forest cover. These cow1ty-by-county notes provide a third unique record ofKenrucky's forest resources, one focused specifically at the presence and abundance of chestnut. Information is available for 51 Kentucky counties, mostly located in the eastern part of the state. During the eight year survey peri­ od, nearly 300 stand assessments were recorded, 250 ofwhich were locat­ ed in the eastern mountains and Knobs region and the rest in the Bluegrass, western Knobs and south-central portions of the state. The use of historic records as a tool for reconstructing previous forest species composition must be approached with caution and their biases rec­ ognized. The reliability ofland deed records, for example, may be hampered by species misidentification or selection bias (Whimey, 1994). Surveyors often

Table 1 A mcrican chestnut a11d forest c0Pe1· estimated during county sttt"Veys co11d11cted b_v USDA forest pathologists between 1924 and 1931. Means followed by the same lcttcl" are not sig11iftcant£v diffen11t as determined by T11kcy's means sepamtton test (a = 0.05).

Chestnut Cover Forest Cover Topography Physiographic Region n= Mean Max Mean Max Median % % 50 20.3 a 50 82.9 a 100 Mountainous 164 6.7 b 50 65.0 b 100 Low Mountains Eastern Knobs 41 5.4 b 20 41 .2 c 90 Hilly Western Knobs 9 2.6 be 20 14.6 d 50 Rolling Mississippian Plateau 8 2.0 be 10 19.0 d 80 Rolling to Hilly Bluegrass 20 0.5 c 2 14.5 d 45 Rolling Total 292 8.2 50 58.4 100

--- 38 JOURNAL OT' IHI· A,\ll RI< AN CHFSTI\ll"I fOUNIHTION science and natural history selected trees based on lifespan, timber value, ease in identification, and size. Land settlement patterns present an additional source of bias, which may alter the interpretation of both the land deed and chestnut blight surveys. Early land settlement was most often concentrated on the lower portions of the landscape with property boundaries corresponding to streanlCourses. Chestnut, on the other hand, occurs most commonly on steep slopes and ridges. In spite ofthese and other shortcomings, few other alternatives exist for estimating the pre-blight abundance of chestnut. Physical evidence of chestnut abundance based on the density of stumps, logs and sprouts are becoming less reliable with time. The rate of log or stump decomposition and of chestnut sprout mortality vary across the landscape and bias abun­ dance estimates that are based on the density of residual material.

RESULTS Based on a 1919 timber inventory, chestnut was listed as a significant por­ tion of the board footage in 75 of Kentucky's 119 counties (Fig. l ). Statewide, chestnut represented 8.5% of the 24,000,000 MBF tallied. Chestnut was distributed throughout the Cumberland Mountains and Plateau, the Knobs Region surrounding the Bluegrass Region, and was scattered throughout the western portion of the state. In 4 counties chest­ nut was the top-ranked species, representing >20% of the 1919 timber inventory. More than 80% of Kentucky's chestnut stumpage occurred in the 27 counties comprising the Cumberland Mountain and Plateau regions. In over half the counties ofthe eastern mountainous counties and in about one-third of the central counties the standing volume of chestnut timber ranked among the top five species. West ofLouisville, chestnut ranked among the rop 5 species in only 2 counties and was top-ranked in none. Chestnut occurred in only 2 of 27 Bluegrass counties and in 3 of 8 Mississippi Embayment counties in extreme western Kentucky. Statewide, white oak (QJ~ercus alba) was the most dominant timber species. It was ranked among the upper five species in 97, 98, and 85% of the counties in the eastern, cen­ tral and western sections of the state, respectively. Similar to American chest­ nut, the abundance ofchestnut oak (Qprinus) reached its maximum \\~thin the Cumberland Mountains and Plateaus and declined westword. Black oak (Q velutina) and hickory (Caryn spp.) both increased west of the Cumberlands while sugar maple (Acer saccharum) and American beech ( Fagus gmndifo­ lia) reached their highest abundance in the central portion of the state.

VOLUME XV, NU~IBER] • SUMMER/FALL 2001 39 science and natural history

American chesmur averaged 8% of the forest cover in the eastern third of Kentucky according to USDA forest pathologists (Table 1). The great­ est abundance of chestnut occurred in the Cumberland Mountain region where chestnut averaged 20% and reached 50% of the forest canopy. Chestnut was significantly scarcer in the less mountainous regions and was nearly absent from the Bluegrass, according to blight survey notes. Counties with the greatest chestnut cover also supported the greatest forest cover; the five counties with tl1e most chestnut were all more than 80% forested. Chestnut corner markers were found on 90% of the land deeds sur­ veyed in the Cumberland Mountain region; they represented 19% of the stems (Table 2 ). Chesmut stems did not exceed 44% within any region. Land deeds in the Bluegrass and western regions did not mention chest­ nut, confirming evidence from the other sources of data.

DISCUSSION The three historic sources estimated the abundance ofchestnut with sur­ prising consistency considering that each source quantified different stand variables (Table 2). The sources each partitioned Kentucky into mree wnes based chesmut

Bluegrass Eastern Knobs Western Knobs Cumberland Plateau Mississippi Embayment

Mississippi Cumberland Plateau Mountains

Chestnut Timber (% of Standing Timber) o 01"2 l.1-S ~. 1 ~ 10 10 I-IS i IS.1-U Figure 1 American chestnut as a percent of the volume of total standing timber stocks in 1919

40 )OURNAL OF THE AMbRICAN Cl IESTNUT FOUNDATION scien ce a nd n atura l h is t ory

Table 2 The pre-blight abundance of American chestnut in Kentucky estimated from independent information sources.

Deeds Records Timber Inventory Bli~ht Survey 1890-1910 1919 1 24-1931 Stems Standing Volume Canopy Cover Physiographic Region Mean Max Mean Max Mean Max % Cumberland Mountains 19 44 17 22 20 so J Cumberla nd Plateau 6 43 7 15 7 50 Eastern Knobs 6 44 7 13 5 20 Western Knobs 2 36 7 19 3 20 Mississippian Plateau 7 40 s 24 2 10 Blueg rass 0 0 0 1 2 Western Coalfields 0 0 1 3 nd nd Mississippi Em bayment 0 0 0 nd nd

abundance. The Cumberland Mountains ofextreme southeastern Kentucky contained the greatest density ofchestnut. The Cumberland Plateau, Knobs and Mississippian Plateau supported intermediate chestnut abundance. Chestnut was relatively rare in the Bluegrass and western extremity of the state. The vast majority of Kentucky's chestnut timber and the highest cover and stem densities were confined to a three-county area in the extreme southeastern corner of the state. Early writings noted the great abundance of chestnut in the Cumberland Mountains·(deFreise, 1884; Braun, 1935 & 1950). The Cumberlands form the western boundary of Braun's Oak-Chestnut fo rest region which fo llows the Blue Ridge and Ridge and Valley physiographic provinces (Fenneman, 1938) and extends northwest from the Tennessee-North Carolina border to central Connecticut (Braun, 1950). This region includes Pine and Cumberland Mountains, Kentucky's only thrust-fault mountains, where chestnut grows under a wide variety ofsoil, topographic and abiotic conditions. On xeric sites \\~th shallow, rocky soils on Pine, Cumberland and Brush Mountains, chestnut was associated \vi th a "stunted growth of hardy trees" including "mountain" chestnut oak, red oak, and various pines (deFreise, 1884). Conversely, on richer sites and deeper soils of Black Mountain, chestnut co-occurred with "yel-

VOLUME XV, NUMBER 1 • SU/\IMFR/FAIL 2001 41 science and natural history

low poplar, black walnut, white and blue ash, birch, linden (basswood), and white hickory (butternut)." A survey ofPine Mountain Braun ( 1935) reported that chestnut was the only overstory species present in all 21 com­ munities situated on both the north- and south-facing slopes on soils formed from shale, sandstone, and limestone parent material. Chestnut was most abundant in Cumberland Mountain oak-chestnut forest (mean: 25% of forest canopy), but was also common in xeric pine-oak stands, mesic mixed mesophytic cove forest, and sugar maple- hemlock- and beech­ dominated stands (Braun 1935, & 1950). Chestnut abundance was intermediate for the majority of Kentucky's eastern and central counties. This region which includes 71 of Kentucky's 120 present-day counties corresponds to the mixed mesophytic forest and oak-hickory forest of the central hardwood region (Bryant, et al. 1993; Hinkle, et al. 1993). The Cumberland Plateau and Knobs of eastern Kenrucky contained only about one-third as much chestnut on average as the Cumberland Mountain region; the mean combined across data types was 6% compared to 19% (Table 2 ). The Mississipian Plateau and western Knobs regions that cover the central third of Kenntcky contained similar or only slightly less chestnut than the Cumberland Plateau area. Each data source, however, noted similar maximum \"alues ofchestnut canopy cover, standing volume, and stem density bet\\'een the second-tier regions and the Cumberland Mountains. It is interesting to note that while the blight surveyors mentioned occasional clumps of pure chestnut, at the stand level, chestnut did not surpass half the canopy cover. Statewide, chestnut represented the highest percent of the standing timber stock (24%) in Larue County, directly south of Louisville on the Mississipian Plateau. Chestnut comprised less than 1% of the forest resources in the remain­ der of Kentucky: the Bluegrass Region and the western-most portion of the state. Our findings agree with original distribution maps regarding the lack of chestnut from the Bluegrass Region (Saucier, 1973; Russell, 1987). Blight sur\'ey notes indicated that chestnut's occurrence within the Bluegrass corresponded to topographic anomalies such as the rocky ledges along the Kentucky Ri\'er, rather than the rolling Inner Bluegrass plain. While the paucity of chestnut in the Bluegrass is often attributed to the underlying limestone parent material, limestone is also the dominant sub­ strate beneath the Mississipian Plateau and is common with the Knobs region (Karathanam, 1992 & 1993) It is unclear if chestnut\ absence

42 JouR.'\AL oF ·rn1 AM~ RJc-1.:-; CH1s1st·1 r<>t' ~1Hnos sci e n ce an d n at u ra l history

from the Bluegrass relates to soil chemical differences between the Ordovician limestone of the Bluegrass and the Mississipian limestone sed­ iments found elsewhere or to some other factor (i.e. high plant compe­ tition, low fire frequency or intensity). Chestnut was found as far west as Carlisle County, bordering the Mississippi River where it represented 0.1 % of the timber inventory. Deeds surveyed around 1900 noted chestnut as a corner tree no farther west than Edmonson and Barren counties near Bowling Green and Mammoth Cave NationaJ Park. Our review of historic records verifies that American chestnut was an important part of Kentucky's pre-blight forests and helps to delimit priori­ ty regions for reintroduction. It remains uncertain how soil and climatic con­ ditions interacted with disturbance processes such as fire, insects, or disease to generate the pattern of chestnut abundance that the early European set­ tlers encountered in Kentucky's forests. As the challenge of reintroducing chestnut to the southern Appalachian region progresses it will be cmcial to consider how enviromnental fuctors and disturbance events may have changed since the time when chestnut was a dominant part of the forest overstory.

\d.nm kd~cmcnt' -The Amencan Chesmut Foundanon and the UK Dcpanmcnt of forestry provided financial suppon for this work. Thanks to Dr. Sandra Anagnostakis for shanng access to USDA forest pathology surYey records, 1he UK Special Collecrions Library for assess to historic deed records, and to Julian Campbell of the Nature Conservancy for helpful ad,·icc and suggestions.

RbFrRENCFS Barton, J. E. 1919. The amount ofstanding timber in Kentucky. The min­ eral and forest resources ofKentucky. The Department of Geology and Forestry, Vol 1. Series 5. Frankfort, KY. Braun, E. L. 1935. The vegetation ofPine Mountain, Kentucky: An analy­ sis of the influence of soils and slope exposure as determined by geo­ logical structure. The American Midland Natttralist 16: 517-565. Braun, E. L. 1950. Deciduous Forests ofEastem North America. Blakiston Co., Philadelphia. Bryant, W. S., W. C. McComb, and J. S. fralish. 1993. Oak-hickory forests (Western mesophytic/Oak hickory forests). Pages 143-201 in W. H. Martin, S. G. Boyce, and A. C. Echternacht, eds. Biodiversity of the southeastern United States: Upland terrestrial communities. John Wiley and Sons, Inc., New York.

Vo1 UMr. xv, NUMBE'R l • SUMMER/FALL 2001 43 science and natural history

Braun, E. L. 1950. Deciditous Forests ofEastern North America. Blakiston Co., Philadelphia. Cochran, M. F. 1990. Chestnuts: Making a comeback. National Geographic 177: 128-140. 1 DeFriese, L. H. 1884. Report on the timbers of the north Cumberland: Bell and Harlan cow1ries. Kentucky Geological Swvey, Frankfurt, KY. pp 81-102. Delcourt, H. R., and P. A. Delcourt. 1997. PreColombian Native American use offire on southern Appalachian landscapes. Conservation Biology 11: 1010-1014. Delcourt, P.A., and H. R. Delcourt. 1998. The influence of prehistoric human-set fires on oak-chestnut forest in the southern Appalachians. Castanea 63: 337-345. Fenneman, N. M. 1938. Physiography ofeastern United States. McGraw Hill, New York. Hebard, F. V., P.H. Sisco, and P.A. Wood. 2000. Meadowview Notes 1999- 2000. The Journal ofthe American Chestnut Foundation 14: 7-15. Hinkle, C.R., W. C. McComb, J.M. Safley Jr., and P.A. Schmalzer. 1993. Mixed mesophytic forests. Pages 203-253 in W. H. Martin, S. G. Boyce, and A. C. Echternacht, eds. Biodiversity ofthe southeastern United States: Upland terrestrial communities. John Wiley and Son, New York. Karathanasis, A. D. 1992. Descriptions and laboratory data for some soils in Kentucky. 6. Knobs Region. University of Kentucky, College of Agriculture, Lexington, KY. pp. 84. Karathanasis, A. D. 199 3. Descriptions and laboratory data for some soils in Kentucky. 7. Eastern Pennyroyal Region. University of Kentucky, College of Agriculture, Lexington, KY. pp. 78 Russell, E. W. B. 1987. Pre-blight distribution of Castanea dentata (Marsh.) Borkh. Bulletin of the Torrey Botanical Club 114: 183-190. Saucier, J. R. 1973. American chestnut ... an American wood. USDA Forest Service. Smalley, G. W. 1986. Classification and evaluation of forest sites on the Northern Cumberland Plateau. USDA For. Serv. pp 74. Whitney, G. G. 1994. From coastal wilderness to fruited plain. Cam bridge University Press, Cambridge. Whi01ey, G. G., and J.P. DeCant. 2001. Government land office surveys and other early land surveys. Pages 147-175 in D. Egan and E. A. Howell, eds. The historical ecology handbook. Island Press, Washington, DC.

44 JOURNAL OF T HE AMERJCAN CH ESTNUT FOUNDATION